Predator identity and time of day interact to shape the risk-reward trade-off for herbivorous coral reef fishes.

Non-consumptive effects (NCEs) of predators occur as prey alters their habitat use and foraging decisions to avoid predation. Although NCEs are recognized as being important across disparate ecosystems, the factors influencing their strength and importance remain poorly understood. Ecological context, such as time of day, predator identity, and prey condition, may modify how prey species perceive and respond to risk, thereby altering NCEs. To investigate how predator identity affects foraging of herbivorous coral reef fishes, we simulated predation risk using fiberglass models of two predator species (grouper Mycteroperca bonaci and barracuda Sphyraena barracuda) with different hunting modes. We quantified how predation risk alters herbivory rates across space (distance from predator) and time (dawn, mid-day, and dusk) to examine how prey reconciles the conflicting demands of avoiding predation vs. foraging. When we averaged the effect of both predators across space and time, they suppressed herbivory similarly. Yet, they altered feeding differently depending on time of day and distance from the model. Although feeding increased strongly with increasing distance from the predators particularly during dawn, we found that the barracuda model suppressed herbivory more strongly than the grouper model during mid-day. We suggest that prey hunger level and differences in predator hunting modes could influence these patterns. Understanding how context mediates NCEs provides insight into the emergent effects of predator-prey interactions on food webs. These insights have broad implications for understanding how anthropogenic alterations to predator abundances can affect the spatial and temporal dynamics of important ecosystem processes.

Reefscapes of fear: predation risk and reef hetero-geneity interact to shape herbivore foraging behaviour.

Predators can exert strong direct and indirect effects on ecological communities by intimidating their prey. The nature of predation risk effects is often context dependent, but in some ecosystems these contingencies are often overlooked. Risk effects are often not uniform across landscapes or among species. Indeed, they can vary widely across gradients of habitat complexity and with different prey escape tactics. These context dependencies may be especially important for ecosystems such as coral reefs that vary widely in habitat complexity and have species-rich predator and prey communities. With field experiments using predator decoys of the black grouper (Mycteroperca bonaci), we investigated how reef complexity interacts with predation risk to affect the foraging behaviour and herbivory rates of large herbivorous fishes (e.g. parrotfishes and surgeonfishes) across four coral reefs in the Florida Keys (USA). In both high and low complexity areas of the reef, we measured how herbivory changed with increasing distance from the predator decoy to examine how herbivorous fishes reconcile the conflicting demands of avoiding predation vs. foraging within a reefscape context. We show that with increasing risk, herbivorous fishes consumed dramatically less food (ca. 90%) but fed at a faster rate when they did feed (ca. 26%). Furthermore, we show that fishes foraging closest to the predator decoy were 40% smaller than those that foraged at further distances. Thus, smaller individuals showed muted response to predation risk compared to their larger counterparts, potentially due to their decreased risk to predation or lower reproductive value (i.e. the asset protection principle). Habitat heterogeneity mediated risk effects differently for different species of herbivores, with predation risk more strongly suppressing herbivore feeding in more complex areas and for individuals at higher risk of predation. Predators appear to create a reefscape of fear that changes the size structure of herbivores towards smaller individuals, increases individual feeding rates, but suppresses overall amounts of primary producers consumed, potentially altering patterns of herbivory, an ecosystem process critical for healthy coral reefs.

Predation risk, resource quality, and reef structural complexity shape territoriality in a coral reef herbivore.

For many species securing territories is important for feeding and reproduction. Factors such as competition, habitat availability, and male characteristics can influence an individual's ability to establish and maintain a territory. The risk of predation can have an important influence on feeding and reproduction; however, few have studied its effect on territoriality. We investigated territoriality in a haremic, polygynous species of coral reef herbivore, Sparisoma aurofrenatum (redband parrotfish), across eight reefs in the Florida Keys National Marine Sanctuary that were either protected or unprotected from fishing of piscivorous fishes. We examined how territory size and quality varied with reef protection status, competition, predation risk, and male size. We then determined how territory size and quality influenced harem size and female size to understand the effect of territoriality on reproductive potential. We found that protected reefs trended towards having more large predatory fishes and that territories there were smaller but had greater algal nutritional quality relative to unprotected reefs. Our data suggest that even though males in protected sites have smaller territories, which support fewer females, they may improve their reproductive potential by choosing nutritionally rich areas, which support larger females. Thus, reef protection appears to shape the trade-off that herbivorous fishes make between territory size and quality. Furthermore, we provide evidence that males in unprotected sites, which are generally less complex than protected sites, choose territories with higher structural complexity, suggesting the importance of this type of habitat for feeding and reproduction in S. aurofrenatum. Our work argues that the loss of corals and the resulting decline in structural complexity, as well as management efforts to protect reefs, could alter the territory dynamics and reproductive potential of important herbivorous fish species.